{"type": "root", "children": [{"type": "p", "children": [{"type": "t", "text": "\nSMAD5 is a receptor‐regulated intracellular mediator that transduces bone morphogenetic protein (BMP) signals, a pathway tightly intertwined with transforming growth factor–β (TGF‐β) signaling. Acting downstream of BMP type I receptors, SMAD5 (together with its close paralog SMAD1) orchestrates a variety of cellular responses including the regulation of proliferation, differentiation, and apoptosis. Its activation is elicited by BMP ligands such as BMP‐2, BMP‐4 and BMP‐6, and its proper function is essential for maintaining tissue homeostasis in diverse cell types—from endothelial and granulosa cells to mesenchymal stem cells."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "1", "end_ref": "3"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn the context of differentiation, SMAD5 plays a pivotal role in promoting osteogenesis. In bone marrow and dental stem cells, its activation via BMP signaling enhances the expression of osteogenic markers such as RUNX2, osteocalcin, and type‐I collagen, thereby stimulating matrix production and cell maturation. SMAD5 also mediates the pro‐osteogenic effects of diverse stimuli—including orthosilicic acid and Genistein—in part by coordinating signals that lead to increased alkaline phosphatase activity and mineralization."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "4", "end_ref": "7"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nBeyond osteogenesis, SMAD5 is integral to the regulation of cell cycle progression and stress responses. In tumor and endothelial cells, for example, mechanical forces such as shear stress elicit SMAD5 activation—often together with SMAD1—to induce G2/M cell cycle arrest, regulate apoptosis and autophagy, and even contribute to premature senescence through modulation of cyclin‐dependent inhibitors. In addition, aberrant regulation of SMAD5 (including its targeting by viral or endogenous microRNAs) may facilitate oncogenic transformation or impact antigenicity in the tumor microenvironment."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "8", "end_ref": "12"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nSMAD5 function is also modulated at the post‐transcriptional level. Its expression in granulosa cells, for instance, is tightly controlled by microRNAs such as miR‐23a, miR‐27a, and miR‐27a‐3p—whose overexpression diminishes SMAD5 levels thereby promoting apoptosis via the FasL‐Fas pathway—while its protein stability is regulated by the E3 ubiquitin ligase Smurf1 that directly interacts with the C terminus of SMAD5 to target it for degradation. Moreover, in fibroblasts and mesenchymal stem cells, the proper SMAD5 signaling axis is essential not only for osteogenic differentiation but also for maintaining extracellular matrix synthesis and proper cellular responses to growth factors."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "13", "end_ref": "19"}]}, {"type": "t", "text": ""}]}, {"type": "t", "text": "\n\n"}, {"type": "p", "children": [{"type": "t", "text": "\nIn addition to its roles in differentiation and cell cycle regulation, SMAD5 is implicated in broader aspects of tissue physiology and disease. In mesenchymal and ovarian cells, for instance, SMAD5 integrates hormonal signals (such as those from follicle-stimulating hormone) with BMP inputs to fine-tune steroidogenesis, while in keloid formation and neural tissues its altered expression has been associated with aberrant tissue remodeling and even cognitive dysfunction in neuropsychiatric conditions. Similarly, in the context of vascular and immune cells, SMAD5 contributes to responses that affect angiogenesis and atherogenesis by balancing pro- and anti-inflammatory signals."}, {"type": "fg", "children": [{"type": "fg_fs", "start_ref": "20", "end_ref": "26"}]}, {"type": "t", "text": ""}]}, {"type": "rg", "children": [{"type": "r", "ref": 1, "children": [{"type": "t", "text": "Drazen B Zimonjic, Marian E Durkin, Catherine L Keck-Waggoner, et al. 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